BIOTROPICA 34(4): 516–533 2002

Distribution and Diversity of Pteridophytes and Melastomataceae

along Edaphic Gradients in Yasunı́ National Park,
Ecuadorian Amazonia1

Hanna Tuomisto 2, Kalle Ruokolainen

Department of Biology, University of Turku, FIN-20014 Turku, Finland
Axel Dalberg Poulsen, Robbin C. Moran 3
Department of Systematic Botany, Aarhus University, Nordlandsvej 68, DK-8240 Risskov, Denmark
Catalina Quintana, Gustavo Cañas, and Jorge Celi
Departamento de Ciencias Biológicas, Herbario QCA, Pontificia Universidad Católica del Ecuador, Av. 12 de
Octubre # 1076 y Carrión, Apartado 17–01–2184, Quito, Ecuador

ABSTRACT
We documented the floristic composition of pteridophytes (ferns and fern allies) and Melastomataceae in Yasunı́
National Park, Amazonian Ecuador. Our main questions were: (1) Are the density of individuals, species richness,
and/or species diversity (measured with Shannon’s H9) of the two plant groups related to edaphic differences? and (2)
How many of the pteridophyte and Melastomataceae species are non-randomly distributed in relation to a soil base
content gradient within terra firme (non-inundated forest). To answer these questions, we sampled 27 line transects
of 500 3 5 m distributed in an area of ca 20 3 25 km. The study area included a permanent 50 ha plot established
to monitor forest dynamics; thus, our results also provide information on landscape-scale floristic variability to which
results from within the plot can be compared. A total of 45,608 individuals and 140 species of pteridophytes, and
4893 individuals and 89 species of the Melastomataceae, were counted in the transects. Both with pteridophytes and
with Melastomataceae, a clear negative correlation was found between the amount of extractable bases in the soil and
the number of plant individuals encountered in a transect. With Melastomataceae, species richness and species diversity
also were negatively correlated with soil base content, but with pteridophytes they were not. More than 50 percent
of the common species of both pteridophytes and Melastomataceae were nonrandomly distributed along the soil cation
content gradient within terra firme. We conclude that while the species richness patterns observed in one plant group
are not indicative of similar patterns in other plant groups, it seems likely that a substantial (but unknown) proportion
of species belonging to other plant groups will be found to show distribution patterns that reflect edaphic preferences
within terra firme forests.

RESUMEN
Se estudió la composición florı́stica de pteridofitas (helechos y plantas afines) y Melastomataceas en el Parque Nacional
Yasunı́, Amazonı́a ecuatoriana. Nuestras preguntas principales fueron: (1) ¿están relacionadas la densidad de individuos,
la riqueza de especies y/o la diversidad (medida con la H9 de Shannon) de los dos grupos de plantas con las diferencias
edáficas? y (2) ¿cuántas de las especies de pteridofitas y Melastomataceas están distribuidas en tierra firme de una
manera que no sea al azar, sino en relación a un gradiente de concentración de cationes en el suelo? Para desarrollar
estas preguntas, muestreamos 27 transectos de 500 por 5 m distribuidos en una área de 20 por 25 km. El área de
estudio incluye una parcela permanente de 50 ha, establecida para estudiar la dinámica del bosque; por lo tanto,
nuestros resultados proveen información sobre la variabilidad florı́stica en una escala del paisaje que puede compararse
con la de la parcela. En total se encontraron en los transectos 45,608 individuos y 140 especies de pteridofitas, y
4893 individuos y 89 especies de Melastomataceas. Tanto las pteridofitas como las Melastomataceas, mostraron una
correlación negativa significativa entre la cantidad de cationes en el suelo y el número de individuos de plantas dentro
de un transecto. En las Melastomataceas, la riqueza de especies y la diversidad estuvieron negativamente correlacionadas
con el contenido de cationes en el suelo, pero en las pteridofitas, no existió tal correlación. Más de 50 por ciento de
las especies comunes de pteridófitas y Melastomataceas estuvieron distribuidas dentro de la tierra firme de una manera
no aleatoria en relación con el gradiente de cationes en el suelo. Concluimos que aunque los patrones de diversidad
de las especies observadas en un grupo de plantas no son indicativos de patrones similares en otros grupos, es probable
que una proporción apreciable (pero desconocida) de especies pertenecientes a otros grupos de plantas pueda mostrar
patrones de distribución que reflejen preferencias edáficas dentro de los bosques de tierra firme.
Key words: Amazonia; diversity; Ecuador; edaphic gradient; Melastomataceae; pteridophytes; species richness; tropical rain
forest.
1 Received 16 December 2001; revision accepted 7 August 2002.
2 e-mail: hantuo@utu.fi
2 Present address: New York Botanical Garden, Bronx, New York 10458-5126, U.S.A.

516

LOCAL PLANT SPECIES RICHNESS (ALPHA DIVERSITY) has
been suggested to vary along edaphic gradients
within the tropical rain forest realm. Extreme hy-
drological conditions, such as seasonal inundation
or permanent waterlogging, have usually been as-
sociated with fewer tree species per unit area than
well-drained conditions in non-inundated or terra
firme areas (Campbell et al. 1986; Balslev et al.
1987; Duivenvoorden 1994, 1996; Clinebell et al.
1995; Duque et al. 2001; Grandez et al. 2001; Ro-
mero-Saltos et al. 2001).
Within the terra firme areas, however, the pat-
terns of species richness variation are still unknown.
Several studies have addressed species richness
trends along a gradient of soil nutrient content, but
the results have been controversial. Some studies
have indicated that species richness increases with
soil fertility (Gentry 1988; Duivenvoorden 1994,
1996; Tuomisto & Poulsen 1996; Tuomisto 1998),
while others have found exactly the opposite (Hus-
ton 1980), a peak at intermediate fertility (Ashton
1992), or no relationship at all (at least if the ex-
tremely poor soil habitats are excluded; Clinebell et
al. 1995).
Another question with no definite answer is
how different plant species are distributed along
edaphic gradients within tropical forests. Some
well-known cases exist of species with a clear dis-
tribution pattern in relation to extreme conditions,
such as permanent waterlogging or extremely nu-
trient-poor soil (e.g., Mauritia flexuosa L.f., which
forms extensive swamp forests in Amazonia, and
the Passiflora vitifolia complex discussed in Gentry
1981).
Plant species distributions in the temperate and
boreal zones are known to be strongly affected by
soil characteristics. Hence to understand species
distribution patterns in tropical forests, it is logical
to study whether or not similar relationships exist
there as well. Few studies, however, have explicitly
documented the distribution of Neotropical plant
species in relation to nutrient contents in the soil.
Duivenvoorden (1995) used canonical correspon-
dence analysis (CCA) to study the distribution of
tree species in relation to drainage, flooding, and
soil nutrient content in an area that covered a wide
range of variability in all of these factors. A simple
classification of pteridophyte species according to
their occurrence in study sites with different soil
cation contents was presented by Tuomisto and
Poulsen (1996). This study was restricted to non-
inundated areas, but it included white sand soils
and hence covered a rather long nutrient gradient.
Pteridophytes and Melastomataceae in Yasunı́ 517
A similarly long gradient was covered in the study
by Tuomisto et al. (1998), which was restricted to
a single fern genus. Often studies dealing with the
responses of plant species to edaphic factors have
used coarse soil classifications (Clark et al. 1998,
1999) rather than directly measured chemical
properties. The question remains, therefore, how
plant species are distributed in areas in which dif-
ferences in soil nutrient contents are less pro-
nounced than in the studies cited above.
The present study was conducted in Yasunı́ Na-
tional Park, Amazonian Ecuador. The study area
included both non-inundated and inundated areas,
of which the non-inundated areas (terra firme) have
traditionally been considered homogeneous (i.e.,
without any edaphically differentiated forest types).
In a parallel paper (Tuomisto et al. in press), we
show that changes in species composition of pte-
ridophytes and Melastomatacae (the latter are pre-
dominantly understory trees, shrubs, and vines) in
the study area are associated with differences in
such factors as the content of exchangeable bases
in the soil, presence or absence of inundation, and
topographic heterogeneity within a transect.
In the present paper, our main questions are
the following: (1) Are the density of individuals,
species richness, and/or species diversity of the two
plant groups related to edaphic differences? and (2)
How many of the pteridophyte and Melastomata-
ceae species are nonrandomly distributed in rela-
tion to the soil base content gradient. The edaphic
gradients considered in the present study are rela-
tively short compared to those in earlier studies.
Thus, instead of representing a special case of ex-
treme landscape-scale variability, our results can be
considered relevant for large areas in western Ama-
zonia, where gradients of this magnitude can be
expected to be common at the landscape to re-
gional scales.
Our results also have importance in another
context. Yasunı́ National Park has recently become
an important study site for tropical botany, with
such long-term research activities as the Forest Dy-
namics Project that is currently inventorying a per-
manent 50 ha tree plot in the area. The first ac-
counts of the floristic composition in subsamples
of the 50 ha plot have already been published (Ro-
moleroux et al. 1997, Svenning 1999, Valencia et
al. in press), and a recent study has inventoried the
woody flora of five 0.1 ha plots in the surroundings
of the 50 ha plot (Romero-Saltos et al. 2001). Our
study provides additional floristic information from
the area.
518 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi
FIGURE 1. Location map of the study area in Ama-
zonian Ecuador. The transects are identified with the
numbers used in the text. The rectangle next to transect
21 is a permanent 50 ha forest dynamics plot.
In the present study, we documented the flo-
ristic composition of pteridophytes and Melasto-
mataceae in the park, using 27 line transects dis-
tributed in an area of ca 20 3 25 km that included
the 50 ha plot. Our objective was to characterize
the overall species composition of the target groups
in the area, which can later be used as a regional
data set against which the results from within the
50 ha plot can be compared.
METHODS
FIELDWORK. Fieldwork was carried out in the vi-
cinity of the Yasunı́ Research Station, situated in
Yasunı́ National Park and operated by Pontificia
Universidad Católica del Ecuador. The sampled
sites were between 08329S and 08469S and from
768219W to 768329W, in an area of ca 25 3 20
km (Fig. 1). The area is accessible by a road con-
structed for oil drilling, and access is presently con-
trolled by the oil company to prevent colonization
and agricultural expansion within the national
park.
Yasunı́ National Park lies 200–300 m above sea
level, and the climate is tropical, humid, and al-
most aseasonal. Annual precipitation exceeds 2400
mm, and no month receives less than 100 mm of
rain on average, although drier months may occur
in some years (Cañadas 1983, Balslev et al. 1987).
Vegetation in the study area consists mainly of
closed-canopy, non-inundated terra firme forests.
The terra firme areas are interrupted by river flood-
plains with seasonally inundated forests and by
swamps dominated by the palm Mauritia flexuosa.
Soil and floristic studies were conducted in 27
sites using transects 500 3 5 m (Fig. 1). The ma-
jority of the transects were placed in non-inundated
forests, but three were placed in seasonally inun-
dated areas for comparison. Transects 6 and 22
were situated in the floodplain of the Tiputini Riv-
er, and transect 13 was situated in the floodplain
of a creek. In the middle of each transect, a narrow
trail (baseline) was opened that followed a prede-
fined compass bearing. For practical reasons, most
transects were started near the road, from which
they were directed deeper into the forest to keep
them in undisturbed vegetation.
The topography of each transect was measured
along the baseline using a clinometer (Suunto, Fin-
land). The difference in elevation between the
highest and lowest point in a transect was used as
a simple measure of topographic heterogeneity,
which was expected to correlate with variation in
moisture availability and other soil differences that
are related to catena formation.
Surface soil samples (the top 5 cm of the min-
eral soil) were collected near the beginning, middle,
and end of each transect. Each of the soil samples
consisted of five pooled subsamples collected with-
in an area of ca 5 3 5 m. Details of the soil analyses
(both methodology and results) will be published
elsewhere (Tuomisto et al. in press). In the present
paper, only information on the total concentration
of extractable bases (Ca, K, Mg, and Na analyzed
in 1 M NH4OAc at pH 7; van Reeuwijk 1993)
will be used.
Pteridophytes (ferns and fern allies) and mem-
bers of the family Melastomataceae (excluding Me-
mecylaceae) were identified to species and counted
within an estimated 2.5 m at both sides of the
baseline for each transect, yielding a sample size of
0.25 ha per transect. To expedite fieldwork, the
lateral limits of the transect were not marked, but
the distance between the baseline and borderline
plant individuals were occasionally measured to
maintain consistency in transect width.
For pteridophytes, only individuals with at least
one leaf longer than 10 cm were recorded, and
epiphytic and climbing individuals were included
only if they had green leaves at a height less than
2 m above ground. Because earlier studies differ in
whether epiphytic pteridophytes were included
(Tuomisto & Ruokolainen 1994, Tuomisto et al.
1995, Ruokolainen et al. 1997) or excluded (Poul-
sen & Balslev 1991; Tuomisto & Poulsen 1996,
2000), we included them but counted ground pte-
ridophytes (self-supporting individuals with rhi-
zomes in the soil) separately from individuals that
were either epiphytic or climbing on a tree trunk

or log. Results will be presented for the data set
containing all individuals, except when explicitly
stated otherwise. For vegetatively spreading species
such as Selaginella, each rooting stem was counted
as one individual.
All individuals of Melastomataceae, including
seedlings that had at least one post-cotyledon leaf,
were recorded. Voucher collections for all species
of both plant groups are deposited in herbaria in
Ecuador (QCA, QCNE), Denmark (AAU), Fin-
land (TUR), and the United States (NY). Unicates
are kept in QCA.
DATA ANALYSES. Chi-square tests were conducted
to determine how many of the pteridophyte and
Melastomataceae species were distributed in a non-
random way across the observed gradient of soil
cation content. Only the 24 terra firme transects
were used in these analyses to avoid the confound-
ing effect of inundation. For the first set of chi-
square analyses, the transects were allocated to two
groups: low and high soil base content (N 5 12 in
both groups; base contents 1.16–6.15 cmol(1)/kg
and 8.97–27.3 cmol(1)/kg, respectively). For the
second set of analyses, transects were allocated to
three groups: low, intermediate, and high soil base
contents (N 5 8 in each group; base contents
1.16–4.81 cmol(1)/kg, 4.89–12.5 cmol(1)/kg,
and 12.7–27.3 cmol(1)/kg, respectively). The ra-
tionale for grouping the transects in two different
ways was that the two-group test allowed identi-
fying species for which occurrences were concen-
trated to either half of the cation gradient, while
the three-group test made it possible to identify
species with occurrences that were concentrated to-
ward the middle of the gradient or to its either
extreme. Only species that occurred in at least 4 of
the 24 transects were tested.
A standard chi-square test assumes that each
plant individual is statistically independent of other
individuals and yields one degree of freedom when
testing whether the corresponding species shows a
nonrandom distribution with respect to the group-
ing criterion. We reasoned that this assumption was
not warranted because individuals within the same
transect were probably highly autocorrelated.
Therefore, we estimated the statistical significance
of each test with a Monte Carlo permutation pro-
cedure using transects as the permutable units in-
stead of individuals: the probability of error of the
observed chi-square statistic was estimated by com-
paring its value to 999 values obtained by permut-
ing transect positions along the cation gradient
while keeping the transect-wise plant abundances
Pteridophytes and Melastomataceae in Yasunı́ 519
fixed. Spatial autocorrelation among transects was
not considered a problem because Mantel tests be-
tween the patterns of geographical and floristic dis-
tances among the transects did not show significant
correlations (Tuomisto et al. in press). The program
Resampling Stats 4.1 was used for the chi-square
tests.
Pearson correlation coefficients were calculated
to test for relationships of individual density, spe-
cies richness, and diversity with soil cation content
and topographic heterogeneity. The correlation co-
efficients were calculated with the program
CorrpPermute (available at http://www.fas.umontreal.
ca/BIOL/Legendre/indexEnglish.html), which allows
estimation of the statistical significance of r with a
Monte Carlo permutation test; 999 permutations were
used.
RESULTS
PTERIDOPHYTES. A total of 45,608 individuals and
140 species of pteridophytes were counted in the
27 transects. Of these, 33,769 individuals (74% of
all individuals) and 108 species (77% of all species)
were found growing on the ground. Because three
species pairs were confused in the field, they have
been lumped for the numerical analyses, which
therefore only deal with 106 species of ground pte-
ridophytes and a total of 137 species (Table 1).
The total number of pteridophyte individuals
found within one transect varied between 734 and
3338, the mean being 1689 individuals per transect
(6760 individuals/ha). The number of pterido-
phyte individuals on the ground varied between
360 and 2521 per transect, the mean being 1251
(5004 individuals/ha). The number of species per
transect varied between 21 and 65 for all pterido-
phytes (x̄ 5 47), and between 12 and 46 for
ground pteridophytes (x̄ 5 35).
The most abundant pteridophyte was Adian-
tum humile Kunze, which was represented by 5000
individuals in 26 transects (Table 1). The other
species with more than 2000 individuals were Poly-
botrya osmundacea Willd., P. crassirhizoma Lellinger,
and Lomariopsis japurensis (Martius) J. Smith. Only
two species were found in all 27 transects, namely
L. japurensis and A. obliquum Willd. Twenty-six
species were found in only one transect, and 11 of
these were represented by a single individual. The
mean number of individuals per species was 330,
and the mean frequency (number of transects in
which the species was found) was 9.
Out of the ten most abundant pteridophyte
species, five were significantly most abundant in
520 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

TABLE 1. Pteridophyte species found in 27 transects in Amazonian Ecuador. A representative voucher specimen is indicated
for each species (collection numbers from the series of R. C. Moran). The general life-form is given for each
species: t 5 terrestrial; sc 5 scandent; cl 5 climber; l 5 lithophyte; and e 5 epiphyte. The total number of
individuals found in each species is shown, as well as the number of terrestrial and scandent individuals as
opposed to epiphytic, epilithic, and climbing individuals. Frequency is the number of transects in which a species
was encountered. Three species pairs were inconsistently identified in the field and are therefore lumped in the
table and in the numerical analyses ( * Antrophyum cajenense and A. guayanense Hieron.; ** Bolbitis
serratifolia and Bolbitis sp. 2; *** Danaea acuminata and D. oblanceolata Stolze).

Number of individuals
Coll. On On other Fre-
Species number Life-form ground substrates Total quency
Adiantum cayennense Klotzsch 6238 t 10 10 5
A. humile Kunze 6237 t 4978 22 5000 26
A. latifolium Lam. 6116 t 11 11 3
A. macrocladum Klotzsch 6256 t 12 12 3
A. cf. olbiquum Willd. 6043 t 1596 12 1608 27
A. petiolatum Desv. 6186 t 22 22 7
A. platyphyllum Sw. 6212 t 3 3 1
A. pulverulentum L. 6159 t 1974 1 1975 26
A. scalare R. M. Tryon 6255 t 1 1 1
A. terminatum Kunze ex Miq. 6050 t 96 1 97 8
A. wilsonii Hook. 6181 t 22 22 1
Adiantum sp. 1 6138 t 19 19 3
Adiantum sp. 2 6239 t, (e) 36 7 43 3
Alsophila cuspidata (Kunze) D. S. Conant 6049 t 209 209 25
Anetium citrifolium (L.) Splitgb. 6082 e 72 72 23
Antrophyum cajenense (Desv.) Sprengel* 6052 e 37 37 15
Asplenium angustum Sw. 6074 e 3 3 1
A. auritum Sw. 6041 e 73 73 11
A. cirrhatum Willd. 6136 e, (t) 76 377 453 22
A. cuneatum Lam. 6268 e 5 5 2
A. juglandifolium Lam. 6103 e, t 4 3 7 4
A. ortegae Murakami and R. C. Moran 6146 t, (e) 862 38 900 18
A. pearcei Baker 6134 e, (t) 5 176 181 22
A. repandulum Kunze 6228 t 1 1 1
A. serratum L. 6269 e 17 17 10
A. stuebelianum Hieron. 6077 e, (t) 59 468 527 22
Asplenium sp. 1 6229 t 87 87 2
Bolbitis lindigii (Mett.) C. Chr. 6046 cl 564 244 808 20
B. nicotianifolia (Sw.) Alston 6112 cl 512 291 803 7
B. oligarchica (Baker) Hennipman 6244 t 542 5 547 15
B. serratifolia (Kaulf.) Schott** 6141 t, l 25 11 36 2
Campyloneurum cf. abruptum (Lindm.)
B. León 6177 e 15 15 3
Campyloneurum aphanophlebium (Kunze)
T. Moore 6185 e 14 14 3
C. cf. brevifolium (Link) Link 6113 e 1 1 1
C. chlorolepis Alston 6114 e 3 3 3
C. fuscosquamatum Lellinger 6065 e 4 726 730 25
C. phyllitidis (L.) C. Presl 6242 e 11 11 10
Campyloneurum sp. 1 6233 t, e 14 7 21 1
Ctenitis refulgens (Mett.) Vareschi 6192 t 2 2 2
Cyathea amazonica R. C. Moran 6045 t 38 38 6
C. lasiosora (Mett. ex Kuhn) Domin 6079 t, (e) 237 17 254 14
C. pungens (Willd.) Domin 6111 t, (e) 380 21 401 20
C. trichiata (Maxon) Domin 6162 t 3 3 2
Cyclodium trianae (Mett.) A. R. Sm. 6047 t 2 2 1
Cyclopeltis semicordata (Sw.) J. Sm. 6132 t 132 132 5
Danaea acuminata H. Tuomisto and
R. C. Moran*** 6209 t 274 274 19
D. bipinnata H. Tuomisto 6260 t 37 37 1
D. falcata H. Tuomisto and 6259 t 23 23 1
R. C. Moran 6064 t 297 1 298 22
 Pteridophytes and Melastomataceae in Yasunı́ 521

TABLE 1. Continued.

Number of individuals
Coll. On On other Fre-
Species number Lifeform ground substrates Total quency
D. nodosa (L.) Sm. 6046 t 297 1 298 22
D. cf. ulei H. Christ 6184 t 70 70 7
Dennstaedtia bipinnata (Cav.) Maxon 6130 t, (e) 7 1 8 4
Didymochlaena truncatula (Sw.) J. Smith 6032 t, (e) 1093 99 1192 25
Diplazium bombonasae Rosenst. 6253 t, (e) 297 7 304 4
D. grandifolium (Sw.) Sw. var.
andicola Stolze 6235 t 83 83 11
D. pinnatifidum Kunze 6029 t 1461 1461 22
D. tungurahuae (Sodiro) C. Chr. 6158 t 734 1 735 14
Elaphoglossum flaccidum (Fée) T. Moore 6227 e 161 161 11
E. luridum (Fée) H. Christ 6179 e 31 31 8
E. cf. megalurum Mickel 6262 e 1 1 1
E. nigrescens (Hooker) Diels 6243 e 45 45 7
E. raywaense (Jenm.) Alston 6153 e 90 90 16
Elaphoglossum sp. 1 6206 e 3 3 1
Hemidictiym marginatum (L.) C. Presl 6163 t, (e) 6 2 8 4
Lindsaea cf. divaricata Klotzsch 6257 t 9 9 3
L. lancea (L.) Bedd. var. lancea 6280 t, (e) 4 4 1
L. phassa K. U. Kramer 6183 t 1 1 1
Lindsaea sp. 1 6211 t 31 31 6
Lomagramma guianensis (Aubl.) Ching 6048 cl 464 610 1074 16
Lomariopsis fendleri D. C. Eaton 6129 e, (cl) 26 26 9
L. japurensis (Mart.) J. Smith 6133 cl 129 1880 2009 27
L. latipinna Stolze 6027 cl 7 505 512 19
L. nigropaleata Holttum 6053 cl 19 789 808 13
Lygodium volubile Sw. 6097 sc 7 7 3
Microgramma fuscopunctata (Hook.)
Vareschi 6071 e 361 361 25
M. percussa (Cav.) Sota 6220 e 2 2 2
Nephrolepis pendula (Raddi) J. Sm. 6083 e, (t) 4 15 19 9
N. rivularis (Vahl.) Mett. ex Krug 6279 e, (t) 30 30 4
Pecluma hygrometrica (Splitg.)
M. G. Price 6247 e 20 20 8
P. ptilodon (Kunze) M. G. Price var.
pilosum (A. M. Evans) Stolze 6246 e 7 7 7
Pityrogramma calomelanos (L.) Link 6273 t 1 1 1
Pleopeltis macrocarpa (Willd.) Kaulf. var.
laciniata Stolze 6161 e 1 1 1
Polybotrya caudata Kunze 6060 cl 1103 554 1657 26
P. crassirhizoma Lellinger 6066 cl 778 1574 2352 26
P. fractiserialis (Baker) J. Sm. 6125 cl 37 5 42 4
P. osmundacea Willd. 6030 cl 1045 1785 2830 25
P. pubens Mart. 6063 cl 105 84 189 5
Polypodium adnatum Klotzsch 6258 e 11 11 1
P. bombycinum Maxon 6037 e 8 8 2
P. caceresii Sodiro 6051 e 1 38 39 14
P. dasypleuron Kunze 6092 e 38 38 11
P. decumanum Willd. 6080 e 8 8 4
Polypodium sp. 1 6119 e 1 1 1
Pteris altissima Poiret 6275 t 2 2 2
P. pungens Willd. 6266 t 4 4 2
Saccoloma inaequale (Kunze) Mett. 6054 t 317 3 320 21
Salpichlaena volubilis (Kaulf.) J. Sm. 6034 sc 126 1 127 17
Selaginella anceps (C. Presl) C. Presl 6170 t, (e) 1109 120 1229 7
S. exaltata (Kunze) Spring 6109 sc 382 382 17
S. geniculata (C. Presl) Spring 6207 t 968 4 972 17
S. haematodes (Kunze) Spring 6223 t, e 65 59 124 7
S. lechleri Hieron. 6276 t 11 11 1
522 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

TABLE 1. Continued.

Number of individuals
Coll. On On other Fre-
Species number Lifeform ground substrates Total quency
S. parkeri (Hook. and Grev.) Spring 6078 t 908 2 910 5
S. revoluta Baker 6101 t 9 9 3
S. speciosa A. Br. 6226 t, (e) 125 53 178 5
Stigmatopteris heterophlebia (Baker)
R. C. Moran 6166 t 227 227 13
S. opaca (Baker) C. Chr. 6126 t 38 38 5
Tectaria cf. antioquoiana (Baker) C. Chr. 6105 t 1911 3 1914 18
T. draconoptera (D. C. Eaton) Copel. 6069 t 96 96 12
T. pilosa (Fée) R. C. Moran 6189 t 56 56 7
T. plantaginea (Jacq.) Maxon 6173 t 33 33 3
T. pubens R. C. Moran 6180 t 169 169 7
T. cf. vivipara Jermy and
T. G. Walker 6190 t 505 2 507 8
Tectaria sp. 1 6148 t 228 228 18
Tectaria sp. 2 6104 t 1252 12 1264 23
Thelypteris abrupta (Desv.) Proctor 6187 t 33 33 9
T. ancyriothrix (Rosenst.)
A. R. Sm. 6215 t 13 13 5
T. arcana (Maxon and C. V. Morton)
C. V. Morton 6198 t 1 1 1
T. biformata (Rosenst.) R. M. Tryon 6107 t, (e) 490 11 501 14
T. dentata (Forssk.) E. St. John 6140 t, e 3 2 5 5
T. glandulosa (Desv.) Proctor
var. brachyodus (Kunze) A. R. Sm. 6218 t 199 199 3
T. hispidula (Decne.) C. F. Reed 6310 t 2 2 1
T. juruensis (C. Chr.) R. M. Tryon and
D. S. Conant 6241 t 838 3 841 6
T. leprieurii (Hook.) R. M. Tryon
var. incana (H. Christ) A. R. Sm. 6312 t 5 5 2
T. macrophylla (Kunze) C. V. Morton 6040 t, (e) 1752 28 1780 15
T. opposita (Vahl) Ching 6311 t 1 1 1
T. opulenta (Kaulf.) Fosberg 6139 e, t 24 39 63 12
T. pennata (Poir.) C. V. Morton 6028 t 483 3 486 17
T. tristis (Kunze) R. M. Tryon 6143 t 11 11 3
Thelypteris sp. 1 6201 t 205 205 3
Thelypteris sp. 2 6142 t 2 2 1
Trichomanes collariatum Bosch 6031 cl 21 21 6
T. diversifrons (Bory) Mett. 6067 t 245 245 21
T. hostmannianum (Klotzsch) Kunze 6087 t 58 58 1
T. pinnatum Hedwig 6264 t 133 133 15
T. polypodiodes L. 6287 e 1 1 1
Trichomanes sp. 1 6265 t 10 10 1
Triplophyllum funestum (Kunze) Holttum 6090 t 55 55 7
137 spp. recognized in numerical analyses 33,769 11,839 45,608
(total 140 spp.)

the transects in which soils had a lower than me-
dian exchangeable base content. Two species
showed a statistically significant bias away from the
poorest soils, while the distribution of three species
was statistically indistinguishable from random
along the soil gradient (Fig. 2). Two of the abun-
dant species, Polybotrya caudata Kunze and L. ja-
purensis, were especially abundant in transects 6
and 22, which were affected by the inundations of
the Tiputini River. Most of the other species were
either rare or absent in these transects. Of course,
even the terra firme transects often included poorly
drained sections in places where the transect
crossed a creek or a poorly drained valley, but as
the present analyses were done at the level of entire
transects, distributions at finer spatial scales were
not assessed.
Out of the 84 pteridophyte species that oc-
 Pteridophytes and Melastomataceae in Yasunı́ 523

FIGURE 2. The abundances (numbers of individuals) of the ten most abundant pteridophyte species (left column)
and the ten most abundant Melastomataceae species (right column) in 27 transects in Amazonian Ecuador. The
transects are arranged in increasing order of soil cation content (sum of exchangeable Ca, K, Mg, and Na). Total
number of individuals of each species is given in parentheses after the species name. Bars corresponding to terra firme
transects are black; those corresponding to transects affected by inundations are gray.

curred in at least 4 of the 24 terra firme transects,
43 (51%) were significantly (P , 0.05) biased in
their distribution among the transect groups that
had been defined according to soil base content
(i.e., they yielded a significant result in the two-
group test or the three-group test, or both). Chi-
square tests gain power when frequencies increase,
but there was no obvious trend in the percentage
of species with biased distributions when the least
frequent species were successively excluded; the
percentage varied but remained between 50 and 60
percent (Fig. 3A).
A clear negative correlation was found between
the amount of extractable bases in the soil and the
524 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

FIGURE 3. Percentage of species of pteridophytes (A) and Melastomataceae (B) that show statistically significant
(chi-square test, P , 0.05) biases in their distribution among two or three groups of non-inundated transects as a
function of the minimum number of transects in which the species were observed. The 24 transects were allocated
to the groups on the basis of the concentration of extractable bases in the soil. The number of species that occurred
in at least four transects is 84 for pteridophytes and 59 for Melastomataceae. The percentages corresponding to the
higher minimum numbers of transects are based on fewer species; the curves are truncated at the point where only
10 species are included. The proportion of species expected to show a statistically significant response by chance is 5
percent both for the two-group test and the three-group test, which is clearly lower than the percentages found here.

number of pteridophyte individuals found in a
transect (r 520.44, P 5 0.03; Fig. 4A). The num-
ber of species and species diversity (as measured
with Shannon’s H9), however, were uncorrelated
with soil base content (Fig. 4C, E). The same re-
sults were obtained whether all 27 transects were
considered, or only the 24 that were situated in
terra firme. When only ground pteridophytes were
included in similar analyses, none of the correla-
tions were statistically significant.
The degree of topographic heterogeneity within
the transect was not significantly correlated with
the number of pteridophyte individuals. The same
result was obtained whether the three seasonally
inundated transects were included in the analysis
or not (Fig. 5A). Both species richness and diversity
were positively correlated with topographic hetero-
geneity, and especially its logarithm, when all tran-
sects were included. This was entirely due to the
three seasonally inundated transects which were
both species-poor and topographically flat; no cor-
relation was found when only terra firme transects
were analyzed (Fig. 5C, E). The results obtained
with ground pteridophytes were similar.
MELASTOMATACEAE. A total of 4893 individuals
and 89 species of the Melastomataceae were count-
ed in the 27 transects (Table 2). The number of
individuals found within one transect varied be-
tween 59 and 372, the mean being 181 individuals
per transect (724 individuals/ha). The number of
species per transect varied between 9 and 44, the
mean being 28.
The most abundant Melastomataceae species
was Miconia triplinervis R. & P., which was repre-
sented by 345 individuals in 18 transects (Table 2).
Miconia serrulata (DC.) Naud. was also represented
by more than 300 individuals. Miconia elata (Sw.)
DC. had the highest frequency as it was found in
23 transects. Eleven species were found in only 1
transect and 3 of these were represented by only 1
individual. The mean number of Melastomataceae
individuals per species was 55, and the mean fre-
quency (number of transects in which the species
was found) was 9.
Out of the ten most abundant Melastomata-
ceae species, five were significantly most abundant
in the transects in which soils had a lower than
median exchangeable base content. Two species
were significantly most abundant in the transects
having soils with a higher base content, while the
distribution of three species was statistically indis-
tinguishable from random along the soil gradient
(Fig. 2). Only one of the abundant species, M.
serrulata, was especially abundant in transects 6 and
22, which were affected by the inundations of the
Tiputini River. The other species were either rare
or absent in these transects.
Out of the 59 Melastomataceae species that oc-
curred in at least four of the terra firme transects,
24 (41%) were significantly biased in their distri-
bution among transect groups defined using soil
 Pteridophytes and Melastomataceae in Yasunı́ 525

FIGURE 4. The number of individuals (A, B), number of species (C, D), and species diversity (as measured with
Shannon’s H9; E, F) plotted against the mean concentration of extractable bases in the soil (sum of Ca, K, Mg, and
Na) in each of 27 transects in Amazonian Ecuador. A, C, E: Pteridophytes; B, D, F: Melastomataceae. Round symbols
indicate terra firme transects; squares indicate transects in river or creek floodplains. The regression line for all 27
transects is shown in black; that for the 24 terra firme transects is shown in gray.

base content (i.e., they yielded a significant result
in the two-group test or the three-group test, or
both). The percentage of species with biased dis-
tributions tended to increase with increasing mi-
numum frequency of the species, until among spe-
cies that had been found in at least ten transects
the percentage more or less stabilized at between
50 and 60 percent (Fig. 3B).
With Melastomataceae, clear negative correla-
tions were found between the amount of extract-
526 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

FIGURE 5. The number of individuals (A, B), number of species (C, D), and species diversity (as measured with
Shannon’s H9; E, F) plotted against the largest elevation difference measured within each of 27 transects in Amazonian
Ecuador. A, C, E: Pteridophytes; B, D, F: Melastomataceae. Round symbols indicate terra firme transects; squares
indicate transects in river or creek floodplains. The best-fitting regression curve for all 27 transects is shown in black;
that for the 24 terra firme transects is shown gray. The nonlinear curves are either logarithmic (C, E) or quadratic (B,
D, F).

able bases in the soil, and the number of individ-
uals, number of species, and Shannon’s H9 diversity
index. The correlation between extractable bases
and individuals was slightly more pronounced
when all 27 transects were considered (r 520.65,
P , 0.001) than when only the 24 terra firme
transects were considered (r 520.62, P , 0.001;
Fig. 4B). The opposite was true of both the cor-
relations between extractable bases and number of
species (r 520.56, P 5 0.004 with all transects, r
520.60, P 5 0.003 with terra firme transects) and
Shannon’s H9 (r 520.46, P 5 0.02 with all tran-
sects, r 520.56, P 5 0.008 with terra firme tran-
sects; Fig. 4D, F).
The degree of topographic heterogeneity within
the transect was negatively correlated with the
number of Melastomataceae individuals when the
seasonally inundated transects were excluded, but
not when they were included in the analysis. Both
with the terra firme data set and the full data set,
a quadratic regression gave a better fit than a
straight line (Fig. 5B). The peak at intermediate
topographic heterogeneity was especially strong
with species richness and diversity, both of which
showed a statistically significant quadratic response
function but no linear correlation. These results
were similar with the full data set and the 24 terra
firme transects (Fig. 5D, F).
COMPARISON OF THE TWO PLANT GROUPS. The Me-
lastomataceae were clearly less abundant than ter-
restrial pteridophytes; the number of Melastoma-
taceae individuals was only 14 percent of the num-
ber of terrestrial pteridophyte individuals. The dif-
ference in species richness, however, was relatively
small; the number of Melastomataceae species was
82 percent of the number of terrestrial pterido-
phyte species.
There was a significant positive correlation be-
tween the number of individuals of the two plant
groups in a transect (r 5 0.64, P 5 0.003), which
became even more pronounced when the three
transects affected by inundations were excluded (r
5 0.72, P , 0.001; Fig. 6A). With all 27 transects,
there was also a positive correlation with the num-
ber of species found in a transect (r 5 0.48, P 5
0.01), but this correlation disappeared when the
three seasonally inundated transects were excluded
(Fig. 6B). The diversity in the transects (as mea-
sured with Shannon’s H9) was not correlated be-
tween the two plant groups no matter whether the
seasonally inundated transects were included or not
(Fig. 6C).
A species’ optimum for soil base content was
Pteridophytes and Melastomataceae in Yasunı́ 527
calculated as the weighted mean of the soil base
contents in the transects where the species was pre-
sent, using species abundance as the weight (Jong-
man et al. 1995). Optima were calculated for all
species with at least ten individuals in the data set,
and they ranged from 1.2 to 19.3 cmol(1)/kg for
pteridophytes and from 3.1 to 19.6 cmol(1)/kg for
Melastomataceae. Although both the minimum
and maximum values were lower for pteridophytes,
the mean was slightly higher [9.1 cmol(1)/kg] than
for Melastomataceae [8.5 cmol(1)/kg]. The fre-
quency distributions of the species optima were
somewhat different between the two plant groups.
Pteridophyte species optima were evenly distribut-
ed along the gradient (although extreme values
were underrepresented), but Melastomataceae spe-
cies optima were strongly skewed toward values be-
low 10 cmol(1)/kg (Fig. 7).
DISCUSSION
The density of pteridophyte individuals varied
widely in the different transects inventoried in Ya-
sunı́: 1440–10,080 individuals/ha for ground pte-
ridophytes and 2940–13,350 individuals/ha for all
pteridophytes. These values span almost the entire
range of 3390 to 13,930 individuals/ha reported
earlier at eight lowland Amazonian sites in Peru
and Ecuador (Tuomisto & Poulsen 1996). Densi-
ties of the Melastomataceae in Yasunı́ ranged from
240 to 1490 individuals/ha, which seems low when
compared to the 1740 individuals/ha reported at a
site in Peru (Tuomisto & Ruokolainen 1994).
Species richness per transect also varied widely
in Yasunı́. The mean species richness (35) of
ground pteridophytes in the Yasunı́ transects is
high compared with quantitative survey samples
from elsewhere in lowland Amazonian Ecuador
and Peru (Tuomisto & Poulsen 1996), which in-
cluded 14–31 species in 0.25 ha. With the Melas-
tomataceae, the species richness values reported
from Peru, 14–22 species/0.35 ha (Tuomisto &
Ruokolainen 1994), fall within the range but below
the mean (28 species/0.25 ha) found in the present
study.
Earlier studies with pteridophytes have shown
that density of individuals, species richness, and di-
versity can vary considerably even at the local scale
(Tuomisto & Poulsen 2000). In some cases, this
variation seemed to be related to topographic var-
iation within a transect, but not always. In the pre-
sent study, the amount of topographic heteroge-
neity within a transect was not correlated with ei-
ther density of individuals, species richness, or di-
528 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

TABLE 2. Melastomataceae species found in 27 transects in Amazonian Ecuador. A representative voucher specimen is
indicated for each species (collection numbers from the series of K. Ruokolainen). The total number of individuals
found in each species is shown. Frequency is the number of transects in which a species was encountered.
Morphospecies numbers are not strictly sequential because the same numbering system is being used in other
study areas, and some of the already numbered morphospecies were not encountered in the present study.

Coll. Number of
number individuals Frequency
Aciotis purpurascens (Aubl.) Tr. 10362 3 1
Adelobotrys boissieriana Cogn. 10353 7 1
Adelobotrys sp. 1 10294 14 6
Adelobotrys sp. 2 10332 49 11
Adelobotrys sp. 4 10380 12 6
Bellucia pentamera Naud. 10357 6 5
Blakea bracteata Gleason 10302 2 2
B. rosea (R. and P.) D. Don 10285 60 21
Clidemia allardii Wurdack 10295 14 7
C. dimorphica Macbr. 10254 139 20
C. epiphytica (Tr.) Cogn. 10355 110 22
C. heterophylla (Desr.) Gleason 10253 92 20
C. japurensis DC. 10390 7 1
C. septuplinervia Cogn. 10261 171 22
C. sprucei Gleason 10340 70 6
Clidemia sp. 3 10323 22 7
Henriettella sp. 1 10344 14 2
Leandra aristigera (Naud.) Cogn. 10378 19 5
L. caquetana Sprague 10319 40 16
L. chaetodon (DC.) Cogn. 10381 6 3
L. longicoma Cogn. 10266 137 22
L. macdanielii Wurdack 10301 11 2
L. secunda (D. Don) Cogn. 10280 15 4
L. secundiflora (DC.) Cogn. 10315 1 1
Leandra sp. 1 10325 10 6
Loreya sp. 1 10333 2 1
Loreya sp. 2 10389 6 4
Maieta guianensis Aubl. 10289 87 13
Miconia abbreviata Markgraf 10268 24 12
M. acutipetala Sprague 10347 17 10
M. amazonica Tr. 10352 1 1
M. ampla Tr. 10379 2 2
M. aurea (D. Don) Naud. 10349 18 9
M. aureoides Cogn. 10368 131 11
M. centrodesma Naud. 10371 7 3
M. decurrens Cogn. 10366 49 11
M. elata (Sw.) DC. 10258 225 23
M. fosteri Wurdack 10270 115 11
M. grandifolia Ule 10278 237 19
M. lamprophylla Tr. 10339 14 3
M. longifolia (Aubl.) DC. 10305 52 5
M. mazanana Macbr. 10312 28 1
M. napoana Wurdack 10343 48 19
M. nervosa (Smith) Tr. 10290 62 15
M. paleacea Cogn. 10296 3 3
M. prasina (Sw.) DC. 10331 13 6
M. procumbens (Gleanson) Wurdack 10252 39 11
M. pterocaulon Tr. 10286 6 3
M. punctata (Desr.) D. Don 10284 96 17
M. schunkei Wurdack 10306 75 10
M. serrulata (DC.) Naud. 10259 307 22
M. tomentosa (L. C. Rich.) D. Don 10276 42 7
M. trinervia (Sw.) D. Don ex Loud. 10264 39 12
M. triplinervis R. and P. 10267 345 18
Miconia sp. 3 10256 80 21
Miconia sp. 4 10320 10 1
 Pteridophytes and Melastomataceae in Yasunı́ 529

TABLE 2. Continued.

Coll. Number of
number individuals Frequency
Miconia sp. 7 10374 109 16
Miconia sp. 12 10334 1 1
Miconia sp. 13 10382 3 2
Miconia sp. 14 10311 9 1
Miconia sp. 15 10372 20 5
Miconia sp. 17 10304 116 7
Miconia sp. 18 10265 234 17
Miconia sp. 19 10375 80 16
Miconia sp. 20 10257 6 3
Miconia sp. 21 10354 21 4
Miconia sp. 22 10338 6 6
Miconia sp. 23 10348 3 2
Miconia sp. 24 10376 213 21
Miconia sp. 25 10346 60 13
Miconia sp. 26 10316 10 3
Miconia sp. 27 10300 14 3
Miconia sp. 29 10321 54 18
Miconia sp. 32 10330 2 1
Miconia sp. 33 10373 48 7
Miconia sp. 34 10281 8 2
Miconia sp. 35 10350 17 2
Miconia sp. 39 10260 2 2
Miconia sp. 40 10318 144 11
Miconia sp. 41 10369 12 5
Miconia sp. 43 10328 19 8
Miconia sp. 44 10282 48 8
Miconia sp. 47 10329 25 5
Monolena primulaeflora Hook. 10335 11 2
Ossaea boliviensis (Cogn.) Gleason 10271 85 13
Tococa caquetana Sprague 10273 103 19
T. discolor Pilger 10360 39 5
Tococa sp. 1 10303 67 3
Triolena amazonica (Pilger) Wurdack 10297 93 11
Total 89 species 4893

versity within the terra firme area. Especially with
species richness, this result is somewhat surprising,
because increasing topographic heterogeneity
would be expected to provide a wider suite of hab-
itats and hence increase beta diversity within the
transect. The expected pattern was found only
when the three transects of floodplain forests were
included in the analysis, but it is likely that this
result was due to the diversity-decreasing effect of
flooding, rather than topography.
With Melastomataceae, very different patterns
were found in relation to topography than with
pteridophytes. A clear peak at intermediate topo-
graphic heterogeneity was found with density of
individuals, species richness, and species diversity.
Not only the flat floodplain transects and some
other relatively flat transects showed low species
richness and diversity, but also the transects with
highest topographic heterogeneity. This surprising
result may be due to the tendency of the transects
with most rugged topography to have relatively
base-rich soils, a factor that correlated negatively
with the species richness and diversity of Melasto-
mataceae. Consequently, the topographic variabili-
ty gradient in the present case is a complex gradient
that not only represents a gradient of heterogeneity
in soil moisture, but where the apparent plant re-
sponse may actually be controlled by different lim-
iting factors in the lower end of the gradient than
in the upper end of the gradient (cf. Pausas & Aus-
tin 2001). Of course, our study has only estab-
lished the existence of correlations and not of caus-
al relationships; to unravel the latter, other soil
properties would have to be included as well, pref-
erably in an experimental framework to clarify the
roles of intercorrelated explanatory factors.
It has often been pointed out that local species
richness is not only dependent on local processes,
530 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

FIGURE 7. Frequency distribution of pteridophyte and

Melastomataceae species optima for different soil base
content intervals in 27 transects in Amazonian Ecuador.
Species represented by less than ten individuals are not
included.

but also regional ones such as immigration from a

metapopulation (Shmida & Wilson 1985, Hubbell
2001). In the present case, the study area was cov-
ered by continuous forest, and even the most dis-
tant transects were not more than 25 km apart; so
it is reasonable to assume that the regional species
pool that was available to colonize a given site was
the same for all transects. Therefore, most of the
variability in species richness among transects was
likely to have depended on factors other than dif-
ferences in the regional species pool or immigration
rate, such as differences in local ecological condi-
tions.
The number of individuals per transect of both
plant groups was negatively correlated with soil
base content, but with species richness and diver-
sity, the trends differed. With Melastomataceae,
species richness and diversity also showed a nega-
tive correlation with soil base content, while with
pteridophytes, there was no correlation. This agrees
with the finding that the two plant groups showed
different trends in the distribution of species opti-
ma for soil base content (the weighted mean soil
base content of sites where each species was found;
Fig. 7). With pteridophytes, the optima were rel-
atively evenly distributed along the soil gradient,
although there was a tendency for intermediate val-
ues to be more common than extreme ones. In
contrast, most Melastomataceae had relatively low
optima for soil base content. Such different distri-
butions probably reflect real differences in the eco-
logical preferences of the two plant groups.
Such contrasting trends in the relationship be-
tween species richness and soil base content be-
tween pteridophytes and Melastomataceae suggest

FIGURE 6. Correlation between pteridophytes and ←

Melastomataceae for number of individuals (A), number that for the 24 terra firme transects is shown in gray.
of species (B), and species diversity (as measured with Symbols indicate the mean concentration of exchangeable
Shannon’s H9; C) in 27 transects in Amazonian Ecuador. bases in the soils as shown in the inset of B.
The regression line for all 27 transects is shown in black;

that species richness of one group of organisms is
not necessarily a very good predictor of species
richness of another group of organisms. This result
agrees with earlier studies conducted both in the
tropics and elsewhere (Prendergast et al. 1993,
Howard et al. 1998). It may be difficult or even
impossible to find a group of organisms that could
serve as a reliable indicator of spatial species rich-
ness patterns in other groups.
The soils in all of our Yasunı́ transects have
relatively high base contents when compared to
other Ecuadorian and Peruvian sites that we have
inventoried. Therefore, the species richness trends
observed here represent only a small part of the soil
variation found in Amazonia, and the patterns ob-
served may not apply to longer edaphic gradients.
In earlier studies, we noticed that in soils with low-
er base contents, both pteridophytes and the Me-
lastomataceae showed a trend of increasing species
richness with increasing soil base content (Tuomis-
to & Ruokolainen 1994, Tuomisto & Poulsen
1996, Tuomisto 1998; K. Ruokolainen, pers. obs.).
These kinds of differences in species richness
trends in different parts of the edaphic gradient
probably explain some of the discrepancies among
the results of earlier studies. Several researchers
have suggested that the response curve of plant spe-
cies richness along a productivity gradient is uni-
modal and attains a peak at intermediate produc-
tivity (Ashton 1992, Rosenzweig & Abramsky
1993). If species richness along a long soil base
content gradient shows this kind of a unimodal
pattern, then the pattern observed along a short
edaphic gradient can be either decreasing, increas-
ing, or flat depending on which part of the long
gradient is sampled (review in Pausas & Austin
2001).
In both pteridophytes and the Melastomata-
ceae, more than half of the common species
showed a statistically significant bias in their abun-
dance distribution in relation to soil base content
(Figs. 2, 3). There is no reason to consider the soil
gradient observed here as exceptionally long in
western Amazonian terms, and therefore, our re-
sults suggest that the majority of pteridophyte and
Melastomataceae species perceive the unflooded
forests of the area as a mosaic of different habitats,
rather than homogeneous. The fact that the species
optima were relatively evenly distributed along the
soil cation gradient, however, indicates that the
habitats need not be sharply differentiated from
each other either floristically or in terms of their
Pteridophytes and Melastomataceae in Yasunı́ 531
environment, but rather that they form a contin-
uum.
Patterns in pteridophyte and Melastomataceae
species composition have been observed to be high-
ly correlated with patterns in tree species compo-
sition (Tuomisto et al. 1995, Ruokolainen et al.
1997, Ruokolainen & Tuomisto 1998, Vormisto et
al. 2000). Therefore, one may expect that not only
pteridophytes and Melastomataceae but also many
tree species, and perhaps species of other plant
groups, show nonrandom distributions across an
edaphic gradient of the length observed in the pre-
sent study. Our results, however, cannot be extrap-
olated to other plant groups without further studies
because big plants may actually show more gener-
alized behavior along edaphic gradients than small
plants (Ruokolainen & Vormisto 2000, Ruokolai-
nen et al. 2002). The tree studies of Clark et al.
(1999) in Costa Rica and Webb and Peart (2000)
on Borneo support our conclusions, while those of
Pitman et al. (1999, 2001) in southern Peru sug-
gest that edaphic differences are less significant.
Out of these studies, however, the Webb and Peart
(2000) study is the only one to include chemical
analyses of soils.
A comparison of species lists (Tables 1 and 2)
and soil properties for Yasunı́ and other study sites
is illuminating. Out of the 49 pteridophyte species
considered rich-soil specialists by Tuomisto and
Poulsen (1996), 45 were found in our Yasunı́ sam-
ple, while out of the 28 intermediate to poor soil
specialists, only 3 were. Similarly, only 4 out of the
24 Melastomataceae species reported by Tuomisto
and Ruokolainen (1994) from a poor to interme-
diate soil site in northern Peru were found in our
present Yasunı́ sample, and all of these seem to be
generalists that have been found also elsewhere in
a wide range of soils (K. Ruokolainen, pers. obs.).
Such results suggest that the observed edaphic pref-
erences of pteridophyte and Melastomataceae spe-
cies are rather consistent even over wider spatial
scales.
ACKNOWLEDGMENTS
We thank Pontificia Universidad Católica del Ecuador for
logistic support, especially its Yasunı́ Research Station and
Herbarium QCA. Renato Valencia is acknowledged for
help in making the logistic arrangements, Oscar Mendoza
and Ermindo Zambrano for assistance in the field work,
and Alan R. Smith and John T. Mickel for help with
species identification. The study was funded by the Dan-
ish Center for Tropical Biodiversity and the Academy of
Finland.
532 Tuomisto, Ruokolainen, Poulsen, Moran, Quintana, Cañas, and Celi

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BIOTROPICA 35(1): 131 2003

ERRATUM

Biotropica 34(4)

IN TABLE 1 [Biotropica 34(4): 520–521 by Tuomisto et al.], the data for D. falcata follow the collection
number 6259. The data following collection number 6064 belong to D. nodosa. The correct collection
number for D. nodosa is 6064 (not 6046).

131